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Ballı Madde Salgısı Honeydew

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Ballı Madde Salgısı Honeydew DOI: http://dx.doi.org/10.16969/teb.67717 Türk. entomol. bült., 2014, 4(2): 121-137 ISSN 2146-975X Derleme (Review) Ballı madde salgısı Honeydew Gordana ĐUROVİĆ1 Selma ÜLGENTÜRK1* Summary Honeydew is excreted (waste) product of Hemipterans Insect related with Sternorrhyncha and Auchenorrhyncha species. Honeydew is liquid syrup substance. Insect diet changes within insect gut facilities, and produce honeydew which contains a variety of saccharides and oligosaccharides also in some proteins in some species. Honeydew has an important positive and/or negative role in particular ecosystems (insect itself which produce honeydew, plant, soil and other organisims). Honeydew has negative impact on plant health due to the formation of fumagine, otherwise positive effects on the nutrient chain. Many microorganisms, insects (such as ants, honey bees, wasps, parasitoids, predators), snails, birds and even man benefits from honeydew as a food. Therefore, honeydew affects these organisms population density and their relationship with each other and the environment. From ancient time, honeydew has been used as a source of food, in the form of manna or honeydew honey by humans. Honeybees is able to utilize honeydew for producing “honeydew honey, forest honey or pine honey”. This honey is an economically important product and widely consumed by humans. Keywords: Honeydew, Sternorrhynca, ants, natural enemies, honeydew honey Özet Ballı madde, Hemiptera takımından genellikle Sternorrhynca olmak üzere, Auchenorrhyncha alttakımına bağlı bazı böcek türlerinin boşaltım (atık) ürünüdür. Şurup kıvamında tatlı bir maddedir. Böcek tarafından emilen bitki özsuyu, sindirim sistemi içinde değiştiğinden, ballı madde, çeşitli sakkaridler ve oligosakkaridler içermektedir. Bazı türlerin ballı maddesinde proteinlerin de olduğu tespit edilmiştir. Ballı madde, ekosistem (salgılayan böcek, bitki, toprak ve diğer canlılar) üzerinde olumlu ve olumsuz çok önemli etkilere sahiptir. Fumajin oluşumu nedeniyle, bitki üzerinde olumsuz etkileri bulunurken, besin döngüsü üzerinde olumlu etkileri bulunmaktadır. Pek çok mikroorganizma, böcekler (karıncalar, balarıları, yaban arıları, parasitoid ve predatörler), kuşlar ve hatta insanlar ballı maddeden besin olarak faydalanmaktadır. Bu nedenle ballı madde, bu canlıların birbiri ve çevresiyle ilişkilerini ve popülasyon yoğunluklarını etkilemektedir. İlk çağlardan beri, insanlar ballı maddeyi bir besin kaynağı olarak, manna ya da salgı balı şeklinde kullanmışlardır. Bal arıları ballı maddeyi toplayarak “Salgı balı, orman balı veya çam balı” üretirler. İnsanlar tarafından severek tüketilen bu bal, ekonomik öneme sahip bir üründür. Anahtar sözcükler: Ballı madde, Sternorrhynca, karınca, doğal düşman, manna, salgı balı 1Ankara Üniversitesi, Ziraat Fakültesi, Bitki Koruma Bölümü, 06110, Dışkapı, Ankara * Sorumlu yazar (Corresponding author) email: [email protected] Alınış (Received): 24.07.2014 Kabul ediliş (Accepted): 21.10.2014 121 Ballı madde salgısı Giriş Ballı madde esas olarak karbonhidrat bakımından zengin, aminoasit ve ikincil bitki bileşenleri bakımından fakir, şurup kıvamında bir maddedir. Bu madde afitler, koşniller, unlubitler, beyazsinekler ve psyllidler gibi Sternorrhynca ve Fulgoridae, Membracidae, Flatidae gibi Auchenorrhyncha (Hemiptera) alt takımlarına bağlı böcek türlerine özgü bir boşaltım (atık) ürünüdür. Floemden beslenen böcekler, vücut hacimlerinin birkaç katı kadar ballı maddeyi kısa sürede vücutlarından atarlar. Emilerek alınan bitki özsuyu, midenin filtre odacığı denilen özel kısmında fazla su ve karbonhidratlar süzülür. Aslında ballı madde, sindirim sisteminde osmotik basıncı düzenleyen mekanizma sonucunda üretilmektedir. Dışarı atılan ballı madde bitkiler, toprak ve insanlarında dahil olduğu geniş bir canlı topluluğunu etkilemektedir. Ballı madde, salgılayan böcek açısından boğulma dahil, çeşitli hijyen ve güvenlik sorunları oluşturabilmektedir (Gullan, 1997). Bitki yüzeyine yayıldığında yumurtadan yeni çıkan nimfler için adeta yapışkan bir tuzaktır. Öte yandan birçok doğal düşman için çekici olmakta, ballı maddeyle beslenmek için gelen doğal düşmanlar, konukçularını da kolaylıkla bulmakta ve avlarıyla beslenmektedir. Ballı madde salgılayan böcekler ile karıncalar arasındaki ilişki en çok çalışılan konudur. Buna ilaveten arılar, kelebekler, hamam böcekleri, salyangozlar, kuşlar ve memelilerin dahil olduğu pek çok canlı ballı madden besin olarak faydalanmaktadır (Camargo & Pedro, 2002; Naskrecki & Nishida, 2007). Balarıları tarafından toplanan ballı madde salgı balı üretiminde kullanılmakta ve bu bal, insanlar tarafından sevilerek tüketilmektedir (Gounari, 2006). Öte yandan ballı madde çok eski çağlardan beri besin olarak (manna, kudret helvası) doğrudan kullanılması, konunun başka bir boyutunu oluşturmaktadır. Bu çalışmada, ballı madde salgılanma mekanizması, salgıyı etkileyen faktörler, ballı madde bileşenleri, doğal düşmanlar üzerindeki etkileri, ballı maddeden besin olarak faydalanan canlılar ve ekositemle olan ilişkileri üzerinde durulmuştur. Ballı Madde salgılayan böceklerde beslenme ve salgılama mekanizması Hemiptera takımına bağlı türlerde ağız parçaları bitkiyi delerek bitki özsuyunu emebilecek şekilde gelişmiştir. Mandibula ve Maxilla’nın iğne şeklinde uzaması ile oluşan iğneler (her birinden ikişer adet), hortum şeklini almış olan labium tarafından korunmaktadır. Bu kılıf (proboscis veya rostrum), coccoidlerde oldukça küçülmüş, hatta körelmiştir. En içte bulunan maxillar stylet’in, karşılıklı duruşları ile iki kanal oluşmuştur. Bunlardan biri bitki özsuyunun emilmesinde, diğeri de tükrük salgılanma görevini almıştır (Gullan & Cranston, 2012). Stiyletler genellikle vücut boyundan uzun olup, ksilem ve/veya floemden bitki özsuyunu emerler (Ben-Dov & Hodgson, 1997). Hemiptera takımında bulunan böceklerin çoğunda sindirim sistemin değişmesiyle filtre odası denilen özel bir kısım meydana gelmiştir (Şekil 1). Bu sistemde farklı yapıda iki boru birbirine yaklaşarak bir doku ile bağlanmıştır. Filtre odasında mesenteron’un iki uzantısı ile protodaeum’un ön kısmı bulunur. Filtre odası, vücut içine alınan fazla miktardaki bitki özsuyunun ve basit şekerlerin hızlı şekilde arka bağırsağa geçirilmesini ve dışarıya atımasını sağlar. Böceklerde ballı madde üretimi, osmotik basıncı ayarlayarak, böceğin iç çevresini ve elektrolitik dengesini düzenlemesine yardımcı olur (Kuran, 1996). 122 Đurović & Ülgentürk, Türk. entomol. bült., 2014, 4 (2) Şekil. 1. Filtre odası (Klowden, 2007). Böceğin floemden aldığı bitki özsuyunun osmotik basıncı, böcek vücudunun osmotik basıncından üç kez daha yüksektir. Bu da böceğin sindirim sistemindeki osmotik basıncı yükseltmekte ve mide duvarından su kaybına yol açmaktadır (Kennedy & Stroyan, 1959; Fisher et al., 1984; Douglas, 2003). Bitki özsuyundaki şekerler, osmotik basınç için çok önemlidir. Alınan disakkarid şekerlerin oligosakkaritlere enzimatik transformasyonu (Fisher et al., 1984; Walters & Mullin, 1988; Rhodes et al.,1997; Wilkinson et al., 1997; Douglas, 2006) ve şekerin asimilasyonu yoluyla böcekler osmotik basınç farkını düzenlemektedir (Mittler & Meikle, 1991). Afitlerdeki monosakkaritlerin glukoz ve fruktoza dönüştürülmesinde glikosidaz enzimi rol oynamakta (Walters & Mullin, 1988) ve ancak bu şekilde mide duvarından geçebilmektedirler (Ashford et al., 2000). Afitler, yüksek sukroz içerikli özsuyu olan bitkilerden beslendiği zaman melezitoz gibi oligosakkaritlerin üretimine geçebilmektedir (Rhodes et al., 1997). Karınca tarafından daha fazla ve sıklıkla ziyaret edilen türlerde tipik olarak daha fazla melezitoz üretilmesi, karıncaların emici böceği daha fazla oranda beslenmeye zorlamasıyla açıklanabilir (Yao & Akimoto, 2001; Woodring et al., 2004). Karley et al. (2005) bezelye afitine (Acyrthosiphon pisum, Harris; Hemiptera: Aphididae) sukroz inhibitörü akarboz enjekte edildikten sonra, 0.75 M´lik sukroz diyetinin ozmotik basıncını düzenlemede başarısız olduğunu göstermiş ve böylece mide ozmolaritesinin düzenlenmesindeki ilke için doğrudan kanıt sağlamışlardır. Son zamanlarda, sukroz (ya da teknik olarak α -glukosidaz) APS1 de bir bezelye afidinden klonlanmış ve enzim, arka-mide membranında gösterilmiştir (Price et al., 2007). Bu da o bölgede gözlenen sukroz aktivitesiyle tutarlılık göstermektedir (Ashford et al., 2000; Cristofoletti et al., 2003). Taşınma muhtemelen şeker taşıyıcısı Ap_ST3´ün (şeker transfer geni) hareketiyle olmaktadır. Ayrıca muhtemelen APS1´in hareketiyle glukoz, transglukozidasyon yolu ile bir sukroz birimine daha sonra bağlanabilir (Ashford et al., 2000;). Böylece melezitoz ya da –aynı prensiple- diğer oligosakkaridleri meydana getirebilirler. Bu oligosakkaritler de daha sonra ballı maddeye ilave edilir (Rhodes et al., 1997; Ashford et al., 2000; Vantaux et al., 2011). Bazı araştırmacılar şeker transformasyonunda mikroorganizmaların etkili olduğunu kaydetmektedir. Bemisia argentifolii (=B. tabaci) Gennadius (Hemiptera: Aleyrodidae)’nin ballı maddesinde bulunan trehaluloz’un, böceğin misetositlerinde bulunan obligat mikroorganizmalar tarafından üretilmiş önemli bir bileşen olduğu düşünülmektedir (Bogo & Mantle, 2000). Bemisia tabaci (Q biyotip) midesinde bulunan Bacillus cinsi bakteriler, oligosakkaridler ve disakkaritlerin sentezinde rol oynamaktadır (Wilkinson et al., 1997). Buchnera aphidicola (Enterobacteriaceae), afitlerde bulunan primer obligat bir bakteridir. Buchnera cinsi bakteriler birkaç aminoasitin sentezinde rol oynarlar (Buchner, 1965; Sasaki et al., 1993; Douglas, 1993; Munson et al., 1991). Yeni bir transkriptomik
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